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      • Record: found
      • Abstract: found
      • Article: found
      Is Open Access

      Minimal information for studies of extracellular vesicles 2018 (MISEV2018): a position statement of the International Society for Extracellular Vesicles and update of the MISEV2014 guidelines

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      Journal of Extracellular Vesicles

      Taylor & Francis

      extracellular vesicles, exosomes, ectosomes, microvesicles, minimal information requirements, guidelines, standardization, microparticles, rigor, reproducibility

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          There is no author summary for this article yet. Authors can add summaries to their articles on ScienceOpen to make them more accessible to a non-specialist audience.

          ABSTRACT

          The last decade has seen a sharp increase in the number of scientific publications describing physiological and pathological functions of extracellular vesicles (EVs), a collective term covering various subtypes of cell-released, membranous structures, called exosomes, microvesicles, microparticles, ectosomes, oncosomes, apoptotic bodies, and many other names. However, specific issues arise when working with these entities, whose size and amount often make them difficult to obtain as relatively pure preparations, and to characterize properly. The International Society for Extracellular Vesicles (ISEV) proposed Minimal Information for Studies of Extracellular Vesicles (“MISEV”) guidelines for the field in 2014. We now update these “MISEV2014” guidelines based on evolution of the collective knowledge in the last four years. An important point to consider is that ascribing a specific function to EVs in general, or to subtypes of EVs, requires reporting of specific information beyond mere description of function in a crude, potentially contaminated, and heterogeneous preparation. For example, claims that exosomes are endowed with exquisite and specific activities remain difficult to support experimentally, given our still limited knowledge of their specific molecular machineries of biogenesis and release, as compared with other biophysically similar EVs. The MISEV2018 guidelines include tables and outlines of suggested protocols and steps to follow to document specific EV-associated functional activities. Finally, a checklist is provided with summaries of key points.

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          Most cited references 315

          • Record: found
          • Abstract: found
          • Article: not found

          Biogenesis, secretion, and intercellular interactions of exosomes and other extracellular vesicles.

          In the 1980s, exosomes were described as vesicles of endosomal origin secreted from reticulocytes. Interest increased around these extracellular vesicles, as they appeared to participate in several cellular processes. Exosomes bear proteins, lipids, and RNAs, mediating intercellular communication between different cell types in the body, and thus affecting normal and pathological conditions. Only recently, scientists acknowledged the difficulty of separating exosomes from other types of extracellular vesicles, which precludes a clear attribution of a particular function to the different types of secreted vesicles. To shed light into this complex but expanding field of science, this review focuses on the definition of exosomes and other secreted extracellular vesicles. Their biogenesis, their secretion, and their subsequent fate are discussed, as their functions rely on these important processes.
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            • Record: found
            • Abstract: found
            • Article: not found

            Argonaute2 complexes carry a population of circulating microRNAs independent of vesicles in human plasma.

            MicroRNAs (miRNAs) circulate in the bloodstream in a highly stable, extracellular form and are being developed as blood-based biomarkers for cancer and other diseases. However, the mechanism underlying their remarkable stability in the RNase-rich environment of blood is not well understood. The current model in the literature posits that circulating miRNAs are protected by encapsulation in membrane-bound vesicles such as exosomes, but this has not been systematically studied. We used differential centrifugation and size-exclusion chromatography as orthogonal approaches to characterize circulating miRNA complexes in human plasma and serum. We found, surprisingly, that the majority of circulating miRNAs cofractionated with protein complexes rather than with vesicles. miRNAs were also sensitive to protease treatment of plasma, indicating that protein complexes protect circulating miRNAs from plasma RNases. Further characterization revealed that Argonaute2 (Ago2), the key effector protein of miRNA-mediated silencing, was present in human plasma and eluted with plasma miRNAs in size-exclusion chromatography. Furthermore, immunoprecipitation of Ago2 from plasma readily recovered non-vesicle-associated plasma miRNAs. The majority of miRNAs studied copurified with the Ago2 ribonucleoprotein complex, but a minority of specific miRNAs associated predominantly with vesicles. Our results reveal two populations of circulating miRNAs and suggest that circulating Ago2 complexes are a mechanism responsible for the stability of plasma miRNAs. Our study has important implications for the development of biomarker approaches based on capture and analysis of circulating miRNAs. In addition, identification of extracellular Ago2-miRNA complexes in plasma raises the possibility that cells release a functional miRNA-induced silencing complex into the circulation.
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              • Record: found
              • Abstract: found
              • Article: not found

              Melanoma exosomes educate bone marrow progenitor cells toward a pro-metastatic phenotype through MET.

              Tumor-derived exosomes are emerging mediators of tumorigenesis. We explored the function of melanoma-derived exosomes in the formation of primary tumors and metastases in mice and human subjects. Exosomes from highly metastatic melanomas increased the metastatic behavior of primary tumors by permanently 'educating' bone marrow progenitors through the receptor tyrosine kinase MET. Melanoma-derived exosomes also induced vascular leakiness at pre-metastatic sites and reprogrammed bone marrow progenitors toward a pro-vasculogenic phenotype that was positive for c-Kit, the receptor tyrosine kinase Tie2 and Met. Reducing Met expression in exosomes diminished the pro-metastatic behavior of bone marrow cells. Notably, MET expression was elevated in circulating CD45(-)C-KIT(low/+)TIE2(+) bone marrow progenitors from individuals with metastatic melanoma. RAB1A, RAB5B, RAB7 and RAB27A, regulators of membrane trafficking and exosome formation, were highly expressed in melanoma cells. Rab27A RNA interference decreased exosome production, preventing bone marrow education and reducing, tumor growth and metastasis. In addition, we identified an exosome-specific melanoma signature with prognostic and therapeutic potential comprised of TYRP2, VLA-4, HSP70, an HSP90 isoform and the MET oncoprotein. Our data show that exosome production, transfer and education of bone marrow cells supports tumor growth and metastasis, has prognostic value and offers promise for new therapeutic directions in the metastatic process.
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                Author and article information

                Journal
                J Extracell Vesicles
                J Extracell Vesicles
                ZJEV
                zjev20
                Journal of Extracellular Vesicles
                Taylor & Francis
                2001-3078
                2018
                23 November 2018
                : 7
                : 1
                Affiliations
                [1 ]A.C.Camargo Cancer Center , São Paulo, Brazil
                [2 ]Aalborg University Hospital, Department of Clinical Biochemistry , Aalborg, Denmark
                [3 ]Aalborg University Hospital, Department of Clinical Immunology , Aalborg, Denmark
                [4 ]Aalborg University, Clinical Institute , Aalborg, Denmark
                [5 ]Aarhus University, Department of Clinical Medicine , Aarhus, Denmark
                [6 ]Aberystwyth University, Institute of Biological, Environmental and Rural Sciences (IBERS) , Aberystwyth, United Kingdom
                [7 ]Academic Medical Centre of the University of Amsterdam, Department of Clinical Chemistry and Vesicle Observation Centre , Amsterdam, The Netherlands
                [8 ]Advanced Center for Chronic Diseases , Santiago, Chile
                [9 ]Aix-Marseille Université, Institut Paoli-Calmettes, INSERM U1068, CNRS UMR7258, Centre de Recherche en Cancérologie de Marseille , Marseille, France
                [10 ]American Red Cross, Scientific Affairs , Gaithersburg, MD, USA
                [11 ]Amsterdam University Medical Centers, Department of Pathology , Amsterdam, The Netherlands
                [12 ]Aston University, School of Life & Health Sciences , Birmingham, UK
                [13 ]AstraZeneca, Discovery Sciences, IMED Biotech Unit , Cambridge, UK
                [14 ]AstraZeneca, Discovery Sciences, IMED Biotech Unit , Gothenburg, Sweden
                [15 ]Atlantic Cancer Research Institute , Moncton, Canada
                [16 ]BC Cancer, Canada’s Michael Smith Genome Sciences Centre , Vancouver, Canada
                [17 ]Beth Israel Deaconess Medical Center , Boston, MA, USA
                [18 ]Boston University School of Medicine , Boston, MA, USA
                [19 ]Brigham and Women’s Hospital, Center for Interdisciplinary Cardiovascular Sciences , Boston, MA, USA
                [20 ]Brown University, Women and Infants Hospital , Providence, RI, USA
                [21 ]Cancer Institute of JFCR , Tokyo, Japan
                [22 ]Cancer Research Institute Ghent , Ghent, Belgium
                [23 ]Cardiff University, School of Medicine , Cardiff, UK
                [24 ]Case Western Reserve University, Department of Medicine , Cleveland, OH, USA
                [25 ]Cedars-Sinai Medical Center , Los Angeles, CA, USA
                [26 ]Cedars-Sinai Medical Center, Smidt Heart Institute , Los Angeles, CA, USA
                [27 ]Center for Psychiatry and Neuroscience, INSERM U894 , Paris, France
                [28 ]Central Research Laboratories, Sysmex Co ., Kobe, Japan
                [29 ]Centro de Biología Molecular Severo Ochoa, Instituto de Investigación Sanitaria la Princesa (IIS-IP) , Madrid, Spain
                [30 ]Children’s Hospital of Los Angeles , Los Angeles, CA, USA
                [31 ]Chinese Academy of Sciences, Wenzhou Institute of Biomaterials and Engineering , Wenzhou, China
                [32 ]CIC bioGUNE, CIBERehd, Exosomes Laboratory & Metabolomics Platform , Derio, Spain
                [33 ]Cincinnati Cancer Center , Cincinnati, OH, USA
                [34 ]City of Hope Comprehensive Cancer Center, Beckman Research Institute, Department of Molecular Medicine , Duarte, CA, USA
                [35 ]Clínica las Condes, Extracellular Vesicles in Personalized Medicine Group , Santiago, Chile
                [36 ]CNC , Coimbra, Portugal
                [37 ]CNR Institute of Neuroscience , Milan, Italy
                [38 ]Copenhagen Lupus and Vasculitis Clinic, Section 4242 - Rigshospitalet , Copenhagen, Denmark
                [39 ]CSGI - Research Center for Colloids and Nanoscience , Florence, Italy
                [40 ]Danube University Krems, Department for Biomedical Research and Christian Doppler Laboratory for Innovative Therapy Approaches in Sepsis , Krems an der Donau, Austria
                [41 ]Department of Veterans Affairs , San Francisco, CA, USA
                [42 ]Duke University Medical Center, Departments of Medicine and Immunology , Durham, NC, USA
                [43 ]Duke University, Department of Ophthalmology , Durham, NC, USA
                [44 ]Durham VAMC, Medical Research Service , Durham, NC, USA
                [45 ]École normale supérieure , Paris, France
                [46 ]Department of Pathology, Erasmus MC, Erasmus Optical Imaging Centre , Rotterdam, The Netherlands
                [47 ]Department of Urology , Rotterdam, The Netherlands
                [48 ]Erasmus MC , Rotterdam, The Netherlands
                [49 ]ETH Zurich, Institute of Pharmaceutical Sciences , Zurich, Switzerland
                [50 ]Eulji University, School of Medicine , Daejeon, South Korea
                [51 ]Evox Therapeutics Limited , Oxford, UK
                [52 ]EVSEARCH.DK , Denmark
                [53 ]Exogenus Therapeutics , Cantanhede, Portugal
                [54 ]Exosomics Siena SpA , Siena, Italy
                [55 ]FAU Erlangen-Nuremberg, Transfusion and Haemostaseology Department , Erlangen, Germany
                [56 ]Federal Research and Clinical Center of Physical-Chemical Medicine , Moscow, Russia
                [57 ]Federal University of Paraná, Department of Genetics, Human Molecular Genetics Laboratory , Curitiba, Brazil
                [58 ]FEMTO-ST Institute, UBFC, CNRS, ENSMM, UTBM , Besançon, France
                [59 ]Finnish Red Cross Blood Service, Research and Development , Helsinki, Finland
                [60 ]Florida State University College of Medicine, Department of Biomedical Sciences , Tallahassee, FL, USA
                [61 ]Fondazione IRCCS Istituto Nazionale dei Tumori, Unit of Immunotherapy of Human Tumors , Milan, Italy
                [62 ]Genome Institute of Singapore, A*STAR , Singapore
                [63 ]Georg-Speyer-Haus Institute for Tumor Biology and Experimental Therapy , Frankfurt, Germany
                [64 ]George Papanicolaou Hospital, Public Cord Blood Bank, Department of Haematology - BMT Unit , Thessaloniki, Greece
                [65 ]Georgetown University, Department of Medicine , Washington, DC, USA
                [66 ]German Armed Forces Central Hospital, Department of General, Visceral and Thoracic Surgery , Koblenz, Germany
                [67 ]German Cancer Consortium (DKTK), Partner Site Freiburg and German Cancer Research Center (DKFZ) , Heidelberg, Germany
                [68 ]German Cancer Research Center (DKFZ), Clinical Cooperation Unit Applied Tumor Biology , Heidelberg, Germany
                [69 ]German Cancer Research Center (DKFZ), Division Signaling and Functional Genomics , Heidelberg, Germany
                [70 ]German Centre for Neurodegenerative Diseases (DZNE) , Bonn, Germany
                [71 ]German Research Center for Environmental Health, Institute for Virology , Munich, Germany
                [72 ]Germans Trias i Pujol Research Institute (IGTP), Can Ruti Campus, REMAR-IVECAT Group , Badalona, Spain
                [73 ]Germans Trias i Pujol University Hospital, Nephrology Service , Badalona, Spain
                [74 ]Ghent University Hospital, Department of Urology , Ghent, Belgium
                [75 ]Ghent University, Department of Radiation Oncology and Experimental Cancer Research, Laboratory of Experimental Cancer Research , Ghent, Belgium
                [76 ]Graduate School of Public Health at the University of Pittsburgh, Division of Occupational and Environmental Medicine , Pittsburgh, PA, USA
                [77 ]Guangzhou Medical University, School of Pharmaceutical Sciences & the Fifth Affiliated Hospital, Key Laboratory of Molecular Target & Clinical Pharmacology , Guangzhou, China
                [78 ]Haaglanden Medical Center, Department of Neurosurgery , The Hague, The Netherlands
                [79 ]Harvard Medical School, Cardiovascular Medicine , Boston, MA, USA
                [80 ]Harvard Medical School, Department of Medicine , Boston, MA, USA
                [81 ]Harvard University, Harvard T.H. Chan School of Public Health , Boston, MA, USA
                [82 ]Harvard University, School of Engineering and Applied Sciences , Cambridge, MA, USA
                [83 ]Hasselt University, Biomedical Research Institute (BIOMED), Department of Medicine and Life Sciences , Hasselt, Belgium
                [84 ]Hasselt University, Biomedical Research Institute (BIOMED) , Hasselt, Belgium
                [85 ]Helmholtz Center Munich German Research Center for Environmental Health, Research Unit Gene Vectors , Munich, Germany
                [86 ]Helmholtz-Centre for Infection Research , Braunschweig, Germany
                [87 ]Helmholtz-Institute for Pharmaceutical Research Saarland , Saarbrücken, Germany
                [88 ]Hiroshima University, Institute of Biomedical & Health Sciences, Department of Cellular and Molecular Biology , Hiroshima, Japan
                [89 ]Houston Methodist Research Institute, Department of Nanomedicine , Houston, TX, USA
                [90 ]Hudson Institute of Medical Research , Melbourne, Australia
                [91 ]Icahn School of Medicine at Mount Sinai, Department of Medicine, Cardiology , New York City, NY, USA
                [92 ]Icahn School of Medicine at Mount Sinai , New York City, NY, USA
                [93 ]IIS-Fundacion Jimenez Diaz-UAM, Department of Nephrology and Hypertension , Madrid, Spain
                [94 ]IKERBASQUE Research Science Foundation , Bilbao, Spain
                [95 ]Imperial College London , London, UK
                [96 ]INEB - Instituto de Engenharia Biomédica , Porto, Portugal
                [97 ]INSERM U1063, Université d’Angers, CHU d’Angers , Angers, France
                [98 ]INSERM U1068, Aix Marseille University, CNRS UMR7258 , Marseille, France
                [99 ]INSERM U1110 , Strasbourg, France
                [100 ]INSERM UMR-S 970, Paris Cardiovascular Research Center , Paris, France
                [101 ]Institució Catalana de Recerca i Estudis Avançats (ICREA) , Barcelona, Spain
                [102 ]Institut Curie, CNRS UMR144, PSL Research University , Paris, France
                [103 ]Institut Curie, INSERM U932, PSL Research University , Paris, France
                [104 ]Institut d’Investigació Germans Trias i Pujol (IGTP), PVREX group , Badalona, Spain
                [105 ]Institut Pasteur de Montevideo, Functional Genomics Unit , Montevideo, Uruguay
                [106 ]Institute of Biomedicine and Molecular Immunology (IBIM), National Research Council (CNR) of Italy , Palermo, Italy
                [107 ]Institute of Medical Biology (IMB), Agency for Science and Technology (A*STAR) , Singapore
                [108 ]Institute of Physiology CAS, Department of Biomaterials and Tissue Engineering, BIOCEV , Vestec, Czech Republic
                [109 ]Institute of Physiology CAS, Department of Biomaterials and Tissue Engineering , Prague, Czech Republic
                [110 ]Instituto Oswaldo Cruz , Rio de Janeiro, Brazil
                [111 ]INSTM - National Interuniversity Consortium of Materials Science and Technology , Florence, Italy
                [112 ]Interuniversity Research Institute for Molecular Recognition and Technological Development (IDM), University of Valencia, Polytechnic University of Valencia , Valencia, Spain
                [113 ]IPATIMUP, Institute of Molecular Pathology and Immunology of the University of Porto , Porto, Portugal
                [114 ]IRCCS - Istituto Ortopedico Rizzoli, Laboratory for Orthopaedic Pathophysiology and Regenerative Medicine , Bologna, Italy
                [115 ]IRCCS Fondazione Don Carlo Gnocchi, Laboratory of Nanomedicine and Clinical Biophotonics (LABION) , Milan, Italy
                [116 ]IRCCS MultiMedica , Milan, Italy
                [117 ]ISGlobal, Hospital Clínic - Universitat de Barcelona, PVREX Group , Barcelona, Spain
                [118 ]Istanbul University Oncology Institute, Basic Oncology Department , Istanbul, Turkey
                [119 ]Jagiellonian University, Faculty of Biochemistry, Biophysics and Biotechnology, Department of Cell Biology , Kraków, Poland
                [120 ]James Cook University, Australian Institute of Tropical Health and Medicine, Centre for Biodiscovery and Molecular Development of Therapeutics , Cairns, Australia
                [121 ]K.G. Jebsen Brain Tumor Research Centre, Department of Biomedicine, University of Bergen , Bergen, Norway
                [122 ]Kansas State University, College of Veterinary Medicine , Manhattan, KS, USA
                [123 ]Karolinska Institute, Clinical Research Center, Department of Laboratory Medicine , Stockholm, Sweden
                [124 ]Karolinska Institute, Clinical Research Center, Unit for Molecular Cell and Gene Therapy Science , Stockholm, Sweden
                [125 ]Karolinska Institute, Department of Medicine and Center for Molecular Medicine, Respiratory Medicine Unit , Stockholm, Sweden
                [126 ]Karolinska Institute, Department of Medicine Solna, Division for Immunology and Allergy , Stockholm, Sweden
                [127 ]Karolinska Institute , Stockholm, Sweden
                [128 ]Kidney Research Centre , Ottawa, Canada
                [129 ]KU Leuven (Leuven University), Department of Human Genetics , Leuven, Belgium
                [130 ]Kyung Hee University, Department of Applied Chemistry , Yongin, Korea
                [131 ]La Trobe University, La Trobe Institute for Molecular Science, Department of Biochemistry and Genetics , Bundoora, Australia
                [132 ]Latvian Biomedical Research and Study Centre , Riga, Latvia
                [133 ]Leibniz Institute for Farm Animal Biology (FBN), Institute of Reproductive Biology , Dummerstorf, Germany
                [134 ]Leiden University Medical Center, Department of Neurosurgery , Leiden, The Netherlands
                [135 ]Loyola University Chicago, Department of Urology , Maywood, IL, USA
                [136 ]Lund University, Department of Cardiology , Lund, Sweden
                [137 ]Maastricht University, GROW, School for Oncology and Developmental Biology, Maastricht Radiation Oncology (MaastRO) Lab , Maastricht, The Netherlands
                [138 ]Maria Sklodowska-Curie Institute - Oncology Center, Gliwice Branch , Gliwice, Poland
                [139 ]Massachusetts General Cancer Center , Boston, MA, USA
                [140 ]Massachusetts General Hospital and Neuroscience Program, Harvard Medical School, Department of Neurology and Radiology , Boston, MA, USA
                [141 ]Massachusetts General Hospital , Boston, MA, USA
                [142 ]Massachusetts General Hospital, Department of Neurology , Boston, MA, USA
                [143 ]Massachusetts General Hospital, Department of Neurosurgery , Boston, MA, USA
                [144 ]Massachusetts General Hospital, Harvard Medical School, Department of Neurology , Boston, MA, USA
                [145 ]Mayo Clinic, College of Medicine, Department of Physiology and Biomedical Engineering , Rochester, MN, USA
                [146 ]Mayo Clinic, Department of Orthopedic Surgery , Rochester, MN, USA
                [147 ]Mayo Clinic, Department of Transplantation Medicine/Department of Physiology and Biomedical Engineering , Jacksonville, FL, USA
                [148 ]Mayo Clinic, Department of Transplantation , Jacksonville, FL, USA
                [149 ]Mayo Clinic , Rochester, MN, USA
                [150 ]McGill University, Division of Experimental Medicine , Montreal, Canada
                [151 ]McGill University , Montreal, Canada
                [152 ]McGill University, The Research Institute of the McGill University Health Centre, Child Health and Human Development Program , Montreal, Canada
                [153 ]McGill University, The Research Institute of the McGill University Health Centre , Montreal, Canada
                [154 ]Medical Center - University of Freiburg, Faculty of Medicine, University of Freiburg, Institute for Infection Prevention and Hospital Epidemiology , Freiburg, Germany
                [155 ]Memorial Sloan Kettering Cancer Center, Department of Medicine , New York City, NY, USA
                [156 ]Monash University , Melbourne, Australia
                [157 ]Mossakowski Medical Research Centre, NeuroRepair Department , Warsaw, Poland
                [158 ]Mount Allison University, Department of Chemistry and Biochemistry , Sackville, Canada
                [159 ]MRC The Gambia , Fajara, The Gambia
                [160 ]MTA-SE Immuno-Proteogenomics Research Groups , Budapest, Hungary
                [161 ]Namur Thrombosis and Hemostasis Center (NTHC), NARILIS , Namur, Belgium
                [162 ]Nanfang Hospital, Southern Medical University, Department of Clinical Laboratory Medicine , Guangzhou, China
                [163 ]National Cancer Center Research Institute, Division of Molecular and Cellular Medicine , Tokyo, Japan
                [164 ]National Cancer Center Research Institute , Tokyo, Japan
                [165 ]National Institute of Chemistry, Department of Molecular Biology and Nanobiotechnology , Ljubljana, Slovenia
                [166 ]National Institutes of Health, National Cancer Institute, Center for Cancer Research , Bethesda, MD, USA
                [167 ]National Institutes of Health, National Cancer Institute, Center for Cancer Research, Laboratory of Pathology , Bethesda, MD, USA
                [168 ]National Institutes of Health, National Institute on Aging , Baltimore, MD, USA
                [169 ]National Institutes of Health, National Institute on Deafness and Other Communication Disorders , Bethesda, MD, USA
                [170 ]National Taiwan University Hospital, Department of Internal Medicine , Taipei, Taiwan
                [171 ]National Tsing Hua University, Department of Power Mechanical Engineering , Hsinchu, Taiwan
                [172 ]National Tsing Hua University, Institute of Nanoengineering and Microsystems , Hsinchu, Taiwan
                [173 ]National University of Singapore, Faculty of Dentistry , Singapore
                [174 ]Nationwide Children’s Hospital , Columbus, OH, USA
                [175 ]Oregon Health & Science University, Department of Anesthesiology & Perioperative Medicine , Portland, OR, USA
                [176 ]Oslo University Hospital Rikshospitalet, Research Institute of Internal Medicine , Oslo, Norway
                [177 ]Oslo University Hospital-The Norwegian Radium Hospital, Institute for Cancer Research, Department of Molecular Cell Biology , Oslo, Norway
                [178 ]Oslo University Hospital, Department of Medical Biochemistry, Blood Cell Research Group , Oslo, Norway
                [179 ]Ospedale Santo Spirito , Pescara, Italy
                [180 ]Ottawa Hospital Research Institute , Ottawa, Canada
                [181 ]Oxford Brookes University, Department of Biological and Medical Sciences , Oxford, UK
                [182 ]Paracelsus Medical University, Department of Transfusion Medicine , Salzburg, Austria
                [183 ]Paracelsus Medical University, GMP Unit , Salzburg, Austria
                [184 ]Philipps University Marburg, Experimental Tumor Biology , Marburg, Germany
                [185 ]Pontificia Universidad Católica de Chile, Advanced Center for Chronic Diseases (ACCDiS) , Santiago, Chile
                [186 ]Pontificia Universidad Católica de Chile, Faculty of Medicine, Department of Hematology-Oncology , Santiago, Chile
                [187 ]POSTECH (Pohang University of Science and Technology), Department of Life Sciences , Pohang, South Korea
                [188 ]Princess Margaret Cancer Centre, University Health Network , Toronto, Canada
                [189 ]QIMR Berghofer Medical Research Institute , Herston, Australia
                [190 ]Queen Mary University of London, Blizard Institute, Epigenetics & Cellular Senescence Group , London, UK
                [191 ]Radboud University Medical Center, Department of Rheumatology , Nijmegen, The Netherlands
                [192 ]Regional Research Network on Extracellular Vesicles, RRNEV , Oslo, Norway
                [193 ]René Rachou Institute/FIOCRUZ , Belo Horizonte, Brazil
                [194 ]Research Service, Olin E. Teague Veterans’ Medical Center , Temple, TX, USA
                [195 ]Royan Institute for Stem Cell Biology and Technology, ACECR, Cell Science Research Center, Department of Stem Cells and Developmental Biology , Tehran, Iran
                [196 ]Saarland University Medical Center, Department of Medicine II , Homburg, Germany
                [197 ]Saarland University , Saarbrücken, Germany
                [198 ]Sanjay Gandhi Postgraduate Institute of Medical Sciences, Department of Molecular Medicine & Biotechnology , Lucknow, India
                [199 ]Sapienza University of Rome, Department of Experimental Medicine , Rome, Italy
                [200 ]Scintillon Institute , La Jolla, CA, USA
                [201 ]Semmelweis University, Department of Genetics, Cell- and Immunobiology , Budapest, Hungary
                [202 ]Semmelweis University, Department of Physiology , Budapest, Hungary
                [203 ]SickKids Hospital, Department of Anesthesia and Pain Medicine , Toronto, Canada
                [204 ]Simon Fraser University, Department of Molecular Biology and Biochemistry , Burnaby, Canada
                [205 ]Sorbonne Université, Centre National de la Recherche Scientifique, Research Unit Biology of Adaptation and Aging (B2A), Team Compensation in Neurodegenerative and Aging (Brain-C) , Paris, France
                [206 ]Spanish Kidney Research Network, REDINREN , Madrid, Spain
                [207 ]Spanish National Cancer Research Center (CNIO), Molecular Oncology Programme, Microenvironment and Metastasis Laboratory , Madrid, Spain
                [208 ]Spinal Cord Injury & Tissue Regeneration Center Salzburg (SCI-TReCS) , Salzburg, Austria
                [209 ]Statens Serum Institut, Department of Autoimmunology and Biomarkers , Copenhagen, Denmark
                [210 ]Stellenbosch University, Department of Physiological Sciences , Stellenbosch, South Africa
                [211 ]Technical University Eindhoven, Faculty Biomedical Technology , Eindhoven, The Netherlands
                [212 ]Technical University of Munich, TUM School of Life Sciences Weihenstephan, Division of Animal Physiology and Immunology , Freising, Germany
                [213 ]Technische Universität Darmstadt, Department of Biology , Darmstadt, Germany
                [214 ]Texas A&M University College of Medicine, Institute for Regenerative Medicine and Department of Molecular and Cellular Medicine , College Station, TX, USA
                [215 ]TGen, Neurogenomics Division , Phoenix, AZ, USA
                [216 ]The Florey Institute of Neuroscience and Mental Health , Melbourne, Australia
                [217 ]The Johns Hopkins University School of Medicine, Department of Molecular and Comparative Pathobiology , Baltimore, MD, USA
                [218 ]The Johns Hopkins University School of Medicine, Department of Neurology , Baltimore, MD, USA
                [219 ]The Johns Hopkins University School of Medicine, Department of Psychiatry and Behavioral Sciences , Baltimore, MD, USA
                [220 ]The Ohio State University, College of Pharmacy, Division of Pharmaceutics and Pharmaceutical Chemistry , Columbus, OH, USA
                [221 ]The Ohio State University , Columbus, OH, USA
                [222 ]The Ohio State University, Comprehensive Cancer Center , Columbus, OH, USA
                [223 ]The School of Biomedical Sciences, University of Western Australia , Perth, Australia
                [224 ]The Scripps Research Institute-Scripps Florida, Department of Molecular Medicine , Jupiter, FL, USA
                [225 ]The Sociedade Beneficente Israelita Brasileira Albert Einstein , São Paulo, Brazil
                [226 ]The University of Melbourne, The Department of Medicine , Melbourne, Australia
                [227 ]The University of Nottingham, School of Medicine, Children’s Brain Tumour Research Centre , Nottingham, UK
                [228 ]The University of Queensland , Brisbane, Australia
                [229 ]The University of Sheffield , Sheffield, UK
                [230 ]The University of Vermont Medical Center, Department of Medicine , Burlington, VT, USA
                [231 ]The Warren Alpert Medical School of Brown University, Department of Medicine , Providence, RI, USA
                [232 ]Thomas Jefferson University, Sidney Kimmel Medical School, Department of Cancer Biology , Philadelphia, PA, USA
                [233 ]Toronto General Hospital Research Institute, University Health Network , Toronto, Canada
                [234 ]TPM of Mirandola , Mirandola, Italy
                [235 ]Trinity College Dublin, School of Pharmacy and Pharmaceutical Sciences, Panoz Institute & Trinity Biomedical Sciences Institute , Dublin, Ireland
                [236 ]Tsinghua University, School of Pharmaceutical Sciences , Beijing, China
                [237 ]UMR 7365 CNRS-Université de Lorraine , Vandœuvre-lès-Nancy, France
                [238 ]UMR-CBMN, CNRS-Université de Bordeaux , Bordeaux, France
                [239 ]UNICAMP, Institute of Biology , Campinas, Brazil
                [240 ]UNIFESP, Departamento de Ciências Farmacêuticas , Diadema, Brazil
                [241 ]Universidad Autónoma de Madrid, Departamento de Biología Molecular , Madrid, Spain
                [242 ]Universidad Autónoma de Madrid, School of Medicine, Department of Medicine , Madrid, Spain
                [243 ]Universidad de Buenos Aires, Facultad de Ciencias Exactas y Naturales, Departamento de Química Biológica , Buenos Aires, Argentina
                [244 ]Universidad de la República, Faculty of Science, Nuclear Research Center, Analytical Biochemistry Unit , Montevideo, Uruguay
                [245 ]Universidade Federal de Paraná , Paraná, Brazil
                [246 ]Universidade Federal de São Paulo Campus Diadema, Departamento de Ciências Farmacêuticas, Laboratório de Imunologia Celular e Bioquímica de Fungos e Protozoários , São Paulo, Brazil
                [247 ]Universidade Federal do Rio de Janeiro, Instituto de Microbiologia , Rio de Janeiro, Brazil
                [248 ]Universidade Federal do Rio Grande do Sul, Instituto de Ciências Básicas da Saúde, Departamento de Microbiologia, Imunologia e Parasitologia , Porto Alegre, Brazil
                [249 ]Università degli Studi di Milano, Department of Clinical Sciences and Community Health, EPIGET LAB , Milan, Italy
                [250 ]Universitat Autònoma de Barcelona, Department of Cell Biology, Physiology & Immunology , Barcelona, Spain
                [251 ]Universitat Autònoma de Barcelona, Hospital Universitari and Health Sciences Research Institute Germans Trias i Pujol, Department of Pathology , Barcelona, Spain
                [252 ]Universitat de València, Departament de Farmàcia i Tecnologia Farmacèutica i Parasitologia, Àrea de Parasitologia , Valencia, Spain
                [253 ]Universitat de València, Health Research Institute La Fe, Joint Research Unit on Endocrinology, Nutrition and Clinical Dietetics , Valencia, Spain
                [254 ]Université Bretagne Loire, Oniris, INRA, IECM , Nantes, France
                [255 ]Université Catholique de Louvain, CHU UCL Namur, Hematology-Hemostasis Laboratory , Yvoir, Belgium
                [256 ]Université Catholique de Louvain, Institut de Recherche Expérimentale et Clinique (IREC), Laboratory of Pediatric Hepatology and Cell Therapy , Brussels, Belgium
                [257 ]Université de Lille, INSERM, U-1192, Laboratoire Protéomique, Réponse Inflammatoire et Spectrométrie de Masse - PRISM , Lille, France
                [258 ]Université de Nantes, INSERM UMR 1238, Bone Sarcoma and Remodeling of Calcified Tissues, PhyOS , Nantes, France
                [259 ]Université de Strasbourg , Strasbourg, France
                [260 ]Université Laval, Centre de Recherche du CHU de Québec, Department of Infectious Diseases and Immunity , Quebec City, Canada
                [261 ]Université Paris Descartes, Sorbonne Paris Cité , Paris, France
                [262 ]Universiteit Hasselt , Diepenbeek, Belgium
                [263 ]University Clinic Eppendorf , Hamburg, Germany
                [264 ]University College London , London, UK
                [265 ]University Hospital Bonn (UKB) , Bonn, Germany
                [266 ]University Hospital Essen, University Duisburg-Essen, Institute for Transfusion Medicine , Essen, Germany
                [267 ]University Hospital Heidelberg, Institute of Pathology, Applied Tumor Biology , Heidelberg, Germany
                [268 ]University Hospital RWTH Aachen, Department of Thoracic and Cardiovascular Surgery , Aachen, Germany
                [269 ]University Hospitals Cleveland Medical Center, Department of Medicine , Cleveland, OH, USA
                [270 ]University Medical Center Göttingen, Developmental Biochemistry , Göttingen, Germany
                [271 ]University Medical Center Göttingen, Hematology and Oncology , Göttingen, Germany
                [272 ]University Medical Center Hamburg-Eppendorf, Department of Neurosurgery , Hamburg, Germany
                [273 ]University Medical Center Hamburg-Eppendorf, Institute of Neuropathology , Hamburg, Germany
                [274 ]University Medical Center Utrecht, Department of Nephrology and Hypertension , Utrecht, The Netherlands
                [275 ]University Medical Center Utrecht, Laboratory for Clinical Chemistry & Hematology , Utrecht, The Netherlands
                [276 ]University of Antwerp, Centre for Proteomics , Antwerp, Belgium
                [277 ]University of Auckland , Auckland, New Zealand
                [278 ]University of Auckland, Department of Molecular Medicine and Pathology , Auckland, New Zealand
                [279 ]University of Auckland, Department of Obstetrics and Gynaecology , Auckland, New Zealand
                [280 ]University of Belgrade, Institute for the Application of Nuclear Energy, INEP , Belgrade, Serbia
                [281 ]University of Birmingham , Birmingham, UK
                [282 ]University of Birmingham, Institute of Microbiology and Infection , Birmingham, UK
                [283 ]University of Brescia, Department of Molecular and Translational Medicine , Brescia, Italy
                [284 ]University of Bristol , Bristol, UK
                [285 ]University of British Columbia Okanagan , Kelowna, Canada
                [286 ]University of Buenos Aires, Instituto de Investigaciones Biomédicas en Retrovirus y SIDA (INBIRS) , Buenos Aires, Argentina
                [287 ]University of California, Davis, Department of Nutrition , Davis, CA, USA
                [288 ]University of California, Davis, Department of Otolaryngology , Davis, CA, USA
                [289 ]University of California, Los Angeles, California NanoSystems Institute , Los Angeles, CA, USA
                [290 ]University of California, Los Angeles, Department of Bioengineering , Los Angeles, CA, USA
                [291 ]University of California, Los Angeles, Department of Pathology and Laboratory Medicine , Los Angeles, CA, USA
                [292 ]University of California, Los Angeles, Jonsson Comprehensive Cancer Center , Los Angeles, CA, USA
                [293 ]University of California, San Diego, Department of Neurosurgery , La Jolla, CA, USA
                [294 ]University of California, San Diego, Department of Obstetrics, Gynecology, and Reproductive Sciences , La Jolla, CA, USA
                [295 ]University of California, San Diego, Department of Pediatrics , San Diego, CA, USA
                [296 ]University of California , San Francisco, CA, USA
                [297 ]University of Cambridge School of Clinical Medicine, Addenbrooke’s Hospital, Department of Medicine, Cambridge NIHR BRC Cell Phenotyping Hub , Cambridge, UK
                [298 ]University of Campinas, Piracicaba Dental School, Department of Oral Diagnosis , Piracicaba, Brazil
                [299 ]University of Chile, Faculty of Chemical and Pharmaceutical Science, Laboratory of Nanobiotechnology and Nanotoxicology , Santiago, Chile
                [300 ]University of Cincinnati College of Medicine , Cincinnati, OH, USA
                [301 ]University of Cologne, Department of Internal Medicine I , Cologne, Germany
                [302 ]University of Colorado, School of Medicine, Department of Ophthalmology, Cell Sight-Ocular Stem Cell and Regeneration Program , Aurora, CO, USA
                [303 ]University of Copenhagen, Faculty of Health and Medical Sciences, Novo Nordisk Foundation Center for Protein Research , Copenhagen, Denmark
                [304 ]University of Copenhagen, Institute of Clinical Medicine , Copenhagen, Denmark
                [305 ]University of Edinburgh, Institute of Immunology & Infection Research , Edinburgh, UK
                [306 ]University of Gothenburg, Institute of Clinical Sciences at Sahlgrenska Academy, Department of Biomaterials , Gothenburg, Sweden
                [307 ]University of Gothenburg, Institute of Clinical Sciences, Department of Surgery, Sahlgrenska Cancer Center , Gothenburg, Sweden
                [308 ]University of Gothenburg, Institute of Medicine at Sahlgrenska Academy, Krefting Research Centre , Gothenburg, Sweden
                [309 ]University of Gothenburg, Sahlgrenska Academy, Department of Rheumatology and Inflammation Research , Gothenburg, Sweden
                [310 ]University of Gothenburg, The Sahlgrenska Academy, Institute of Neuroscience and Physiology, Department of Psychiatry and Neurochemistry , Mölndal, Sweden
                [311 ]University of Helsinki, EV Core Facility , Helsinki, Finland
                [312 ]University of Helsinki, Faculty of Biological and Environmental Sciences, Molecular and Integrative Biosciences Research Programme, EV group , Helsinki, Finland
                [313 ]University of Hertfordshire, School of Life and Medical Sciences, Biosciences Research Group , Hatfield, UK
                [314 ]University of Kentucky, College of Medicine, Department of Physiology , Lexington, KY, USA
                [315 ]University of L’Aquila, Department of Life, Health and Environmental Sciences , L’Aquila, Italy
                [316 ]University of Liège, GIGA-R(MBD), PSI Laboratory , Liège, Belgium
                [317 ]University of Ljubljana, Faculty of Medicine, Institute of Biochemistry , Ljubljana, Slovenia
                [318 ]University of Lyon, INRA, EPHE, UMR754 Viral Infections and Comparative Pathology , Lyon, France
                [319 ]University of Lyon, Lyon-Sud Faculty of Medicine, CarMeN Laboratory (UMR INSERM 1060-INRA 1397) , Pierre-Bénite, France
                [320 ]University of Mainz, Institute of Developmental Biology and Neurobiology , Mainz, Germany
                [321 ]University of Malta, Department of Pathology , Msida, Malta
                [322 ]University of Manchester, Division of Cancer Sciences, Manchester Cancer Research Centre , Manchester, UK
                [323 ]University of Manchester , Manchester, UK
                [324 ]University of Maryland, Fischell Department of Bioengineering , College Park, MD, USA
                [325 ]University of Massachusetts Medical School, RNA Therapeutics Institute , Worcester, MA, USA
                [326 ]University of Michigan Medical School , Ann Arbor, MI, USA
                [327 ]University of Michigan, Biointerfaces Institute , Ann Arbor, MI, USA
                [328 ]University of Michigan, Department of Biomedical Engineering , Ann Arbor, MI, USA
                [329 ]University of Michigan, Department of Internal Medicine - Hematology/Oncology Division , Ann Arbor, MI, USA
                [330 ]University of Michigan, Department of Medicine , Ann Arbor, MI, USA
                [331 ]University of Michigan, Department of Urology , Ann Arbor, MI, USA
                [332 ]University of Minnesota Medical School, Institute on the Biology of Aging and Metabolism, Department of Biochemistry, Molecular Biology and Biophysics , Minneapolis, MN, USA
                [333 ]University of Modena and Reggio Emilia, Division of Oncology , Modena, Italy
                [334 ]University of Nebraska Medical Center, Department of Pharmacology and Experimental Neuroscience , Omaha, NE, USA
                [335 ]University of Notre Dame, Department of Biological Sciences , Notre Dame, IN, USA
                [336 ]University of Oslo, Institute of Clinical Medicine , Oslo, Norway
                [337 ]University of Ottawa , Ottawa, Canada
                [338 ]University of Oulu, Faculty of Medicine, Cancer and Translational Medicine Research Unit , Oulu, Finland
                [339 ]University of Oxford, Department of Physiology, Anatomy and Genetics , Oxford, UK
                [340 ]University of Oxford, Radcliffe Department of Medicine, Acute Stroke Programme - Investigative Medicine , Oxford, UK
                [341 ]University of Padova, Department of Comparative Biomedicine and Food Science , Padova, Italy
                [342 ]University of Padova, Department of Women’s and Children’s Health , Padova, Italy
                [343 ]University of Palermo, Department of Biopathology and Medical Biotechnologies , Palermo, Italy
                [344 ]University of Perugia, Department of Chemistry, Biology and Biotechnology , Perugia, Italy
                [345 ]University of Pisa, Centro Dipartimentale di Biologia Cellulare Cardio-Respiratoria , Pisa, Italy
                [346 ]University of Porto, Faculty of Pharmacy (FFUP), IBMC/I3S , Porto, Portugal
                [347 ]University of Porto, Faculty of Pharmacy (FFUP) , Porto, Portugal
                [348 ]University of Porto, i3S-Instituto de Investigação e Inovação em Saúde , Porto, Portugal
                [349 ]University of Porto, ICBAS - Instituto de Ciências Biomédicas Abel Salazar , Porto, Portugal
                [350 ]University of Rochester , Rochester, NY, USA
                [351 ]University of São Paulo, Ribeirão Preto Medical School, Department of Pathology and Forensic Medicine , Ribeirão Preto, Brazil
                [352 ]University of São Paulo, Ribeirão Preto Medical School , Ribeirão Preto, Brazil
                [353 ]University of Science and Culture, ACECR, Department of Developmental Biology , Tehran, Iran
                [354 ]University of South Alabama, Department of Pharmacology, Center for Lung Biology , Mobile, AL, USA
                [355 ]University of Southern California Keck School of Medicine , Los Angeles, CA, USA
                [356 ]University of Southern California , Los Angeles, CA, USA
                [357 ]University of Technology Sydney, Discipline of Pharmacy, Graduate School of Health , Sydney, Australia
                [358 ]University of Texas MD Anderson Cancer Center, Department of Cancer Biology, Metastasis Research Center , Houston, TX, USA
                [359 ]University of Torino, Department of Medical Sciences , Torino, Italy
                [360 ]University of Torino, Department of Molecular Biotechnology and Health Sciences , Torino, Italy
                [361 ]University of Toronto, Department of Anesthesia , Toronto, Canada
                [362 ]University of Toronto, Department of Laboratory Medicine and Pathobiology , Toronto, Canada
                [363 ]University of Toronto, Department of Medical Biophysics , Toronto, Canada
                [364 ]University of Toronto, Department of Medicine, Division of Neurology , Toronto, Canada
                [365 ]University of Tsukuba , Tsukuba, Japan
                [366 ]University of Virginia Health System, Department of Medicine, Division of Nephrology , Charlottesville, VA, USA
                [367 ]University of Virginia, Flow Cytometry Core, School of Medicine , Charlottesville, VA, USA
                [368 ]University of Würzburg, Rudolf Virchow Center , Würzburg, Germany
                [369 ]Utrecht University, Faculty of Veterinary Medicine, Department of Biochemistry and Cell Biology , Utrecht, The Netherlands
                [370 ]Utrecht University, University Medical Center Utrecht, Center for Molecular Medicine & Regenerative Medicine Center , Utrecht, The Netherlands
                [371 ]Utrecht University, University Medical Center Utrecht, Department of Neurosurgery, Brain Center Rudolf Magnus, Institute of Neurosciences , Utrecht, The Netherlands
                [372 ]Utrecht University, University Medical Center Utrecht, Department of Pathology , Utrecht, The Netherlands
                [373 ]Vanderbilt University Medical Center, Epithelial Biology Center, Department of Medicine , Nashville, TN, USA
                [374 ]Vanderbilt University School of Medicine, Department of Cell and Developmental Biology , Nashville, TN, USA
                [375 ]Veterans Affairs Medical Center , San Francisco, CA, USA
                [376 ]Vilnius University, Institute of Biomedical Sciences, Department of Physiology, Biochemistry, Microbiology and Laboratory Medicine , Vilnius, Lithuania
                [377 ]Vlaamse Instelling voor Technologisch Onderzoek (VITO) , Mol, Belgium
                [378 ]Washington University , Saint Louis, MO, USA
                [379 ]Weill Cornell Medicine, Department of Medicine , New York City, NY, USA
                [380 ]Weizmann Institute of Science, Department of Biomolecular Sciences , Rehovot, Israel
                [381 ]West Virginia University, Department of Chemical and Biomedical Engineering and WVU Cancer Institute , Morgantown, WV, USA
                [382 ]West Virginia University, Department of Microbiology Immunology and Cell Biology , Morgantown, WV, USA
                [383 ]West Virginia University , Morgantown, WV, USA
                [384 ]Xiamen University, Department of Chemical Biology , Xiamen, China
                Author notes
                CONTACT Clotilde Théry Clotilde.Thery@ 123456curie.fr Institut Curie/INSERM U932, 26 rue d’Ulm, 75005Paris, France
                Kenneth W Witwer kwitwer1@ 123456jhmi.edu 733 North Broadway, MRB 829 Baltimore, MD21205, USA
                [*]

                These authors are co-corresponding authors and contributed equally to this work. All authors, except for the corresponding authors, are listed alphabetically.

                This article was first drafted begining in late 2017 and finalized in September, 2018 with several rounds of feedback from the ISEV Executive Board and JEV Editorial Board.

                [&]

                The following authors are members of the 2016-2018 and/or 2018-2020 ISEV Board of Directors: Cherie Blenkiron, Edit I Buzas, Dolores Di Vizio, Uta Erdbrügger, Juan M Falcón-Pérez, Chris Gardiner, Andrew F Hill (current President), Raghu Kalluri, Jan Lötvall (past President), Malene Møller Jørgensen, Rienk Nieuwland, Lorraine O’Driscoll, Susmita Sahoo, Carolina Soekmadji, Hidetoshi Tahara, Ana Claudia Torrecilhas, Marca HM Wauben, Alissa Weaver, Kenneth W Witwer, Hang (Hubert) Yin, Lei Zheng.

                [£]

                The following authors are Editors of the Journal of Extracellular Vesicles: Editors-in-Chief: Clotilde Théry, Peter Quesenberry, Yong Song Gho; Associate Editors: Elena Aikawa, Chantal Boulanger, Edit I Buzas, Giovanni Camussi, Emanuele Cocucci, Dolores Di Vizio, Andrew F Hill, Harry Holthofer, Kwang Pyo Kim, Eva-Maria Krämer-Albers, Saara Laitinen, Cecilia Lässer, Suresh Mathivanan, Esther Nolte-’t Hoen, Takahiro Ochiya, Hector Peinado, Janusz Rak, Neta Regev-Rudzki, Raymond Schiffelers, Marca HM Wauben, Kenneth W Witwer.

                [#]

                Authors who are members of the International Society for Cell and Gene Therapy (ISCT) Exosomes Scientific Committee include Sai Kiang Lim (Co-Chair), Bernd Giebel (Co-Chair), Luis A Ortiz, Donald G Phinney and Daniel J Weiss (ISCT Chief Scientific Officer). Massimo Dominici is Past President of ISCT.

                [$]

                Authors who are members of the joined ISEV-ISAC (International Society for Advancement of Cytometry)-ISTH (International Society on Thrombosis and Haemostasis) EV Flow Cytometry Working group are: Alain Brisson, Frank AW Coumans, André Görgens, Jennifer C Jones, Joanne Lannigan, Sten Libregts, François Mullier, Rienk Nieuwland, John P Nolan, Marca HM Wauben, Joshua A Welsh.

                Article
                1535750
                10.1080/20013078.2018.1535750
                6322352
                © 2018 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group on behalf of The International Society for Extracellular Vesicles.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial License ( http://creativecommons.org/licenses/by-nc/4.0/), which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

                Counts
                Tables: 4, References: 320, Pages: 44
                Product
                Funding
                Funded by: Agence Nationale de la Recherche 10.13039/501100001665
                Award ID: ANR-11-LABX-0043
                Funded by: Agence Nationale de la Recherche 10.13039/501100001665
                Award ID: ANR-10-IDEX-0001-02 PSL
                Funded by: National Institute on Aging 10.13039/100000049
                Award ID: AG057430
                Funded by: National Institute on Drug Abuse 10.13039/100000026
                Award ID: DA040385
                Funded by: INSERM, Institut Curie, Ministry of Education
                Award ID: NA
                Funded by: INCa
                Award ID: INCA-11548
                Funded by: Fondation ARC
                Award ID: PGA1 RF20180206962
                Funded by: Fondation ARC
                Award ID: PJA 20171206453
                Funded by: Sidaction 10.13039/100009060
                Award ID: 17-1-AAE-1138
                Funding is listed only for the corresponding authors. CT’s laboratory is funded by INSERM, Institut Curie, Ministry of Education, and grants from INCa (INCA-11548), French National Research Agency (ANR-10-IDEX-0001-02 PSL* and ANR-11-LABX-0043), SIDACTION (17-1-AAE-1138), Fondation ARC (PGA1 RF20180206962, PJA 20171206453). KWW and CT receive support from NIDA (DA040385). KWW is also supported in part by NIA AG057430, NIDA DA047807 and NIMH MH118164.
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